A Case Study on the Phylogeny and Conservation of Saker Falcon
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Ornis Hungarica 2014. 22(1): 1–14. DOI: 10.2478/orhu-2014-0007 A case study on the phylogeny and conservation of Saker Falcon KRISZTIÁN POMICHAL1*, BALÁZS VÁGI2 & Tibor Csörgő1 Krisztián Pomichal, Balázs Vági & Tibor Csörgő 2014. A case study on the phylogeny and conser- vation of Saker Falcon. – Ornis Hungarica 22(1): 1–14. Abstract The Saker Falcon is a falconid raptor species with Palearctic distribution. It has ne ver been a common bird in Hungary, now there are cc. 220–230 nesting pairs within the count ry borders. Currently total world population is cc. 19 000–34 000 individuals. Its taxonomic status is complica ted. Two subspecies are distinguished (Falco c. cherrug and Falco c. milvipes); however, molecular data does not support this split. Phylogeny of the species is also not clarified, similarly to closely related raptors. There are ma- ny factors threatening the population of the Saker. One of these factors is the occurrence of the hybrid falcons. By molecular investigations more data can be gained, that could be useful in practical conservation, too. Keywords: Falco cherrug, phylogenetics, taxonomy, hybridization, conservation Összefoglalás A kerecsensólyom a sólyomfélék családjába tartozó, palearktikus elterjedésű faj. Magyarországon sosem volt gyakori madár, jelenleg 220–230 pár fészkelhet az országban. A világállomány körülbelül 19 000– 34 000 párt tesz ki. A faj taxonómiája nagyon összetett. Jelenleg két alfaját különböztetik meg, molekuláris bizo- nyítékok azonban nem támasztják alá ezt a felosztást. A faj leszármazási vonala szintén nem tisztázott, ugyanúgy, ahogy a legközelebbi rokon fajoké sem. Számos tényező veszélyezteti a faj állományait, ezek közül az egyik leg- fenyegetőbb a hibrid madarak problematikája. A molekuláris vizsgálatok segítségével olyan ismeretek szerezhe- tőek, melyek haszonnal alkalmazhatóak a gyakorlati természetvédelemben is. Kulcsszavak: Falco cherrug, filogenetika, taxonómia, hibridizáció, természetvédelem 1 Department of Anatomy, Cell- and Developmental Biology, Eötvös Loránd University, 1117 Budapest, Pázmány Péter sétány 1/c, Hungary, e-mail: [email protected] 2 MTA-ATK-NÖVI Lendület Evolutionary Ecology Research Group, 1022 Budapest, Herman Ottó út 15., Hungary *corresponding author Molecular approaches have been gaining in- e.g. previously undivided world populations creasing importance in every field of biolo- could be split into two or more species. This gy in the last 60 years since the description could change conservation categories when of the molecular structure of the DNA. Inves- marginal or isolated subpopulations which tigations on nuclear and mitochondrial ge- comprise only a fraction of the world popu- nomes yielded significant knowledge in seve lation turn out to be separate species. ral streams of biology, which questioned the Certain species may have evolved by hyb re le vance of traditional taxonomic categories ridization and this could be important not and this way affected other research fields as only on a theoretical base but it could have well, e.g. practical nature conservation. practical implications as well. In practice of On one hand, clarifying taxonomic sta- the last decades new hybrids of greater size, tus and relations of recent species could be robustness and superior hunting skills have fascinating, on the other hand new results been created between falconid species. It may challenge widely accepted taxonomy, is dubious whether these hybrids escaping 2 ORNIS HUNGARICA 2014. 22(1) to the wild can reproduce with wild speci- of this was that time common nest looting mens; whether they are capable of creating which concerned mainly nests built on cliffs. viable offspring, and if so how they affect In several cases nesting failure was caused the gene pool of wild populations. by human disturbance, and in certain occa- It is also problematic that there is diffe sion illegal shooting also happened. The spe- rent legislation affecting trade in hybrids cies has been preserved due to enormous ef- originating from breeding stocks. This po forts of nature conservation experts. In 22 ses a threat that captured wild specimens years the population grew from 8 breeding can be traded as hybrids to thwart regula- pairs to 113–145 pairs (Bagyura et al. 2004). tions. Here, we present a case study of such According to latest reports the Hungarian problematic on the Saker Falcon. populations can be estimated 220–230 pairs (Bagyura et al. 2012). Distribution and population size Taxonomy of the Saker Falcon The Saker Falcon (Falco cherrug Gray, 1834) is a palearctic species belonging to The Saker Falcon was described by the Bri the falconid family (Falconidae). The wes tish zoologist John Edward Gray in Hard- tern border of its distribution area is in Cent wicke’s book Illustrations of Indian Zoo ral Europe, while its eastern border is near logy in 1834. The description was made on the Amur River (Ragyov et al. 2010). The a young raptor wintering in India. As resear total area of its nesting range and habitats is chers were not aware of its complete distri- cc. 10 300 square kilometres (BirdLife In- bution area, the Saker was described inde- ternational 2012). The world population of pendently by several taxonomists, thus, its the species is decreasing in its whole range, nomenclature has been confused (Ragyov et the loss was 32% according to 1991–2010 al. 2009). Previously, six subspecies of the estimation means (the exact value should questionable species were described as fol- be somewhere between 30 and 49 per- lows: F. c. cyanopus (distributed in Central cent). Currently total world population of Europe and along the Volga river), F. c. cher- the species is cc. 19 200–34 000 individu- rug (Russia), F. c. hendersoni (Pamir Moun- als (BirdLife International 2012), but there tains, Himalaya), F. c. altaicus (AltaiSayan are also world population size estimates Region), F. c. milvipes (Kyrgyzstan and of 7200–8520 individuals from the year of Mongolia), F. c. coatsi (from Iran to Tian 2003 (Bagyura et al. 2007). Shan) (Nittinger et al. 2007). Some authors’ Saker Falcon has never been a common work (e.g. Pfeffer 2009) also mention the Si- bird in Hungary; but in the 1970s its po berian Saker (Falco c. saceroides) and the pulation reached a critical state. At the begin- Anatolian Saker (F. c. anatolicus). Falco ch. ning of the following decade only 8 breeding saceroides is known as hybrid between the attempts were recorded in the 13 known ter- Gyrfalcon and the Saker, while F. c. anatoli- ritories. Two of them were successful with cus is an other subspecies. 22 fledged juveniles. In 1980 the population Most debated is the population of Central on country level have not been more than 30 Asian mountains (Altai, Tian Shan) which is pairs (Bagyura et al. 2004). The main cause considered to be a separate species (the Al- K. Pomichal, B. Vági & T. Csörgő 3 tai Falcon – Falco altaicus) (Moseikin & El- Phylogenetics lis 2004). Others claim that the Altai Falcon is a persistent hybrid between the Gyrfalcon The first molecular investigations from the and the Saker Falcon. Based on genetic re- 1960s to the middle of the 1980s were based search the Altai Falcon cannot be separated on allozyme variance. Allozymes are dif- from the other two subspecies of the Saker ferent variants of enzymes which are co (Wink et al. 2004). ded at the same locus of the chromosomes. The Saker Falcon is a polytypic species While researchers managed to get informa- with two accepted subspecies: F. c. cherrug tion by direct inspection of the DNA this and F. c. milvipes (Clements et al. 2011). method lost its significance; it has been out- While F. c. cherrug is distributed in Europe competed by DNA sequencing with restric- as well as east from the Yenisei River and tion enzymes and later direct sequencing of the Altai Mountains to China, F. c. milvipes the DNA. Recently the whole genome of the is restricted to the northern and eastern parts Saker Falcon has been sequenced. of Central Asia and NorthEastern China Certain sources treat the Hierofalco group (FergusonLees & Christie 2001). This se as a subgenus (Wink & Seibold 1996), paration is based on phenotypic traits solely. while others consider it only as collective Molecular investigations based on micro- terminus without any taxonomic relevance satellites and mitochondrial control regions (Nittinger et al. 2007). Recent species of of DNA samples extracted from skin and the Hierofalco group originate from Afri- plumage of museum specimens failed to de- ca and they spread out of that continent in tect any differences between ssp. cherrug several consecutive waves. This group con- and ssp. milvipes (Nittinger et al. 2007). sists of four species: the Saker Falcon, the Taxonomy of the species is still concer Gyrfalcon (F. rusticolus), the Laggar Fal- ning researchers. Some taxonomists claim con (F. jugger) and the Lanner Falcon (F. that division into two subspecies is over- biarmicus) (Nittinger et al. 2007). Based on simplified (Karyakin 2009, Pfander 2011). some research the Black Falcon (F. subni- Pfeffer (2009) propose a separation of fur- ger) from Australia is also belonging to this ther six subspecies, one of which (F. c. ana- group (Wink et al. 2004). tolicus) has been still undescribed. This The northern and southern populations concept is based on the species’ undoubted- had been separated from each other by the ly remarkable phenotypic variance, which formation and expansion of the Sahara De- is underpinned by the authors’ observa- sert. Northern populations begun to spread tions by traditional methods in nature and towards the Mediterranean area and colo- captivity; however, it contradicts current nized the whole Eurasian continent in three phylogene tic results (which are otherwise waves during the warmer periods of the cited in that paper). Pleistocene. The Laggar Falcon evolved Pfander (2011) suggests the introduction from population spreading towards India of a currently unused taxonomic concept, through the eastern part of the Mediterrane- the ‘semispecies’ to the taxonomy of the an Basin, while the Gyrfalcon evolved from Saker Falcon.